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Zain, Moss & El-Sheikh<br />

ARTICLE<br />

The structural cell membrane was contiguous with the<br />

membrane of the septum, between the pseudophialide<br />

and merosporangium cross-walls. Here the pseudophialide<br />

necks were cylindrical, 1.2–1.4 µm long, ca. 1–1.1 µm<br />

diam, with a septum delimiting the merosporangium<br />

(Fig. 1E). Pseudophialides from which merosporangia<br />

had been released had a flared-shape structure from the<br />

merosporangium cell wall attached to the pseudophialide<br />

neck; the septum and septal plug, and structure within the<br />

pseudophialide, was shrivelled (Fig. 1J).<br />

Merosporangia and merosporangiospores<br />

Immature merosporangia were obovoid, whereas mature<br />

ones were obovate. Merosporangia matured first on those<br />

pseudophialides towards the “apex” of the sporocladium with<br />

those on the peripheral pseudophialides the last to mature.<br />

The merosporangia of L. pennispora were obovate, 3–4 µm<br />

wide near the base, narrowed distally to 2.0–2.5 µm wide,<br />

18–23 µm long, and possessed regular surface annulation<br />

with interconnecting ridges, but lacked any rod-shaped<br />

ornamentation (Fig. 1G).<br />

The merosporangia were produced terminally or<br />

subterminally and singly on the pseudophialides. A septum<br />

formed at the apex of the pseudophialide neck delimited the<br />

merosporangium from the pseudophialide. Merosporangia<br />

had a three-layered wall continuous with the pseudophialide<br />

wall. The merosporangium wall comprised an outer, 150–<br />

200 nm thick, electron-opaque layer; a middle, 200–250<br />

nm thick, electron-opaque layer; and an inner, 40–50 nm<br />

thick, electron-dense layer (Fig. 1H). Each merosporangium<br />

contained a single merosporangiospore.<br />

The merosporangiospore was obovate, 2.3–2.5 µm diam,<br />

with a ca. 1 µm diam papilla-like base. The merosporangiospore<br />

had a four-layered wall: an outer, 2–5 nm thick, electrondense<br />

layer; adpressed to the outer layer, a thick, 5–10<br />

nm, electron-dense layer; and an innermost fourth, 90–<br />

100 nm thick, amorphous, electron-transparent layer.<br />

The merosporangiospore wall was contiguous with the<br />

merosporangium wall at the distal region, and separated by<br />

an electron-opaque layer at the base of the merosporangium.<br />

Detachment of the merosporangia from their pseudophialides<br />

occurred at the base of the merosporangium.<br />

Detached merosporangia possessed a single, 3–5 µm<br />

long, “appendage” that was attached to the base of the<br />

merosporangium (Fig. 1I). The appendage was acicular,<br />

unbranched, ca. 0.1 µm diam and attached to the<br />

merosporangiospore inner cell wall layer and passed through<br />

the septum plug to the pseudophialide. Pseudophialides from<br />

which merosporangia had been released possessed a flared<br />

collar-like distal region with the septum, which delimited the<br />

pseudophialide from the merosporangium, retained at the<br />

base of the collar (Fig. 1J).<br />

The detachment of the merosporangium from the<br />

pseudophialide occurred by rupture of the merosporangium<br />

wall near the base, when the merosporangiospore wall<br />

becomes coated with the electron-opaque layer (Fig. 1K).<br />

When the merosporangium has been detached, a part<br />

of the merosporangium wall remained attached to the<br />

pseudophialide neck, as well as the septum between the<br />

pseudophialide and merosporangium. The cytoplasmic<br />

layer between the merosporangiospore base and the<br />

merosporangium wall appeared as a spherical-shape<br />

structure attached to the septal-plug.<br />

DISCUSSION<br />

Aerial hyphae of Linderina pennispora were investigated<br />

previously at the ultrastructure level (Young 1969, 1970b,<br />

Benny & Aldrich 1975). The hyphae had a two-layered wall<br />

which comprised an outer amorphous layer and an inner<br />

fibrillar layer (Young 1969, 1970b, Benny & Aldrich 1975).<br />

Numerous spines were described as attached to the outer<br />

layer of the hyphal wall (Young 1970b). Benny & Aldrich<br />

stated that the spines were attached to the inner layer of the<br />

wall and appeared to be covered by material from the outer<br />

wall layer (Benny & Aldrich 1975). The results presented here<br />

show that the surface ornamentations of the aerial hyphae<br />

of Linderina pennispora appears rod-shaped rather than<br />

spine-like. The ornamentation is attached to the outer layer<br />

of the hyphal wall. It is fibrillar, electron-dense, and seem<br />

to be derived from the same material as the outer layer.<br />

There was no definite description for the sporangiophore in<br />

all the previously published studies on the morphology of<br />

the species. This study revealed that the sporangiophore<br />

of L. pennispora arose as a lateral branch of the vegetative<br />

hyphae. The sporangiophores are narrower in diameter<br />

than the vegetative hyphae, and the ontogeny of the<br />

sporangiophore and its sympodial growth are described here<br />

for the first time, and explain the diagnostic “zigzag” form of<br />

the sporangiophore.<br />

Benjamin (1966) described the sporocladia of Kickxellaceae<br />

species as the most highly developed sporiferous<br />

branchlets in Zygomycetes. The present study provides details<br />

of the sporocladia and their ontogeny. The sporocladium<br />

initials are produced terminally by the sporangiophore, and,<br />

when the sporangiophore resumes its growth, the sporocladia<br />

are displaced laterally. The terminal sporocladium is displaced<br />

after the formation of the merosporangia, particularly at<br />

the late stages of merosporangia development. Benny &<br />

Aldrich (1975) observed the surface ornamentations of<br />

pseudophialides of L. pennispora and stated that they were<br />

coated with fewer rod-shape surface ornamentations than<br />

the sporocladium, and comprised a structure they termed<br />

an “abscission vacuole”. However, Young (1974) described<br />

a similar structure in the pseudophialides of Kickxella<br />

alabastrina, and then termed the structure a “labyrinthiform<br />

organelle” based on its morphology. A similar structure was<br />

also demonstrated in the pseudophialides of Dipsacomyces<br />

acuminosporus and Martensiomyces pterosporus (Young<br />

1968). Benny & Aldrich (1975) suggested that this structure<br />

was related to the abscission and dispersal mechanism of<br />

the wet-spored species of Kickxellaceae. They believed that<br />

this structure was produced from the septum delimiting the<br />

merosporangium (Benny & Aldrich 1975).<br />

Our electron microscopic studies of the pseudophialides<br />

of Linderina pennispora show, for the first time, a concentric<br />

arrangement of the pseudophialides on the sporocladium,<br />

and that only the peripheral pseudophialides are coated<br />

with a rod-shaped ornamentation. Ultra-thin sections<br />

106 <br />

ima fUNGUS

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